doi:10.

1093/scan/nsr090 SCAN (2013) 8, 285^299

The social evaluation of faces: a meta-analysis

of functional neuroimaging studies
Peter Mende-Siedlecki,1 Christopher P. Said,2 and Alexander Todorov1,3
1
Department of Psychology, Princeton University, Princeton, NJ, USA, 2Department of Psychology and Center for Neural Science, New York
University, New York, NY, USA, and 3Department of Psychology, Behavioural Science Institute, Radboud University, Nijmegen,
The Netherlands

Neuroscience research on the social evaluation of faces has accumulated over the last decade, yielding divergent results.
We used a meta-analytic technique, multi-level kernel density analysis (MKDA), to analyze 29 neuroimaging studies on face
evaluation. Across negative face evaluations, we observed the most consistent activations in bilateral amygdala. Across positive
face evaluations, we observed the most consistent activations in medial prefrontal cortex, pregenual anterior cingulate cortex
(pgACC), medial orbitofrontal cortex (mOFC), left caudate and nucleus accumbens (NAcc). Based on additional analyses compar-
ing linear and non-linear responses, we propose a ventral/dorsal dissociation within the amygdala, wherein separate populations
of neurons code for face valence and intensity, respectively. Finally, we argue that some of the differences between studies are
attributable to differences in the typicality of face stimuli. Specifically, extremely attractive faces are more likely to elicit
responses in NAcc/caudate and mOFC.

Keywords: meta-analysis; fMRI; social cognition; face perception; face evaluation; trait judgments; trustworthiness; attractiveness

INTRODUCTION personality have been shown to predict important outcomes

Within a single glance of a face, people automatically ap-
praise face attractiveness and make a host of social attribu-
tions (Olson and Marshuetz, 2005; Bar et al., 2006; Willis
and Todorov, 2006; Rule et al., 2009; Todorov et al., 2009).
For example, 33-ms exposure to a face is sufficient for people
to make trustworthiness decisions (Todorov et al., 2009).
Additional time exposure simply increases confidence in
these decisions (Willis and Todorov, 2006). As one of the
founding fathers of modern social psychology, Solomon
Asch (1948, p. 258), put it, ‘We look at a person and imme-
diately a certain impression of his character forms itself in
us. A glance, a few spoken words are sufficient to tell us a
story about a highly complex matter. We know that such
impressions form with remarkable rapidity and with great
ease. Subsequent observations may enrich or upset our view,
but we can no more prevent its rapid growth than we can
avoid perceiving a given visual object or hearing a melody’.
Recent research confirms Asch’s insights (Zebrowitz, 1999;
Todorov et al., 2008a,b; Zebrowitz and Montepare, 2008).
People rapidly and effortlessly form impressions from facial
appearance. Although the validity of such impressions is low
(Olivola and Todorov, 2010a), inferences of character and
Received 21 January 2011; Accepted 27 November 2011
Advance Access publication 27 January 2012
We thank Hedy Kober and Bryan Denny for assistance with the MKDA toolbox, Tor Wager for making the
MKDA toolbox available online, and Jenny Porter and Katie Hansen, who provided the behavioral data for
Figure 5B. We also thank Jamil Zaki, Adam Hampshire, and Leigh Nystrom for their assistance in obtaining
analyses not reported in their original papers. This research was supported by National Science Foundation
grant BCS-0823749 (to A.T.), National Science Foundation grant DGE 1148900 (to P.M.-S.), and the Russell
Sage Foundation.
Correspondence should be addressed to Peter Mende-Siedlecki. E-mail: pmende@princeton.edu
ß The Author (2012). Published by Oxford University Press. For Permissions, please email: journals.permissions@oup.com
in domains ranging from politics (Todorov et al., 2005;
Ballew and Todorov, 2007; Olivola and Todorov, 2010b),
law (Zebrowitz and McDonald, 1991; Blair et al., 2004;
Eberhardt et al., 2006), mate choice (Olivola et al., unpub-
lished data), business (Hamermesh and Biddle, 1994; Rule
and Ambady, 2008) and the military (Mazur et al., 1984).
Despite the importance of first impressions for social
interactions, research on their neural basis is in its infancy.
Researchers began to use social neuroscience methods to
investigate this basis only a decade ago (Adolphs et al.,
1998; Nakamura et al., 1998; Aharon et al., 2001; Winston
et al., 2002). Although a number of neuroimaging studies
have been published on the topic, many of the results have
been inconsistent (Todorov et al., 2011). The objective of
this article is to provide a quantitative summary of the
major findings across studies on face evaluation.
The neural basis of face evaluation
Neuroimaging research on the social evaluation of faces has
usually focused on evaluations along the trait dimensions of
trustworthiness and attractiveness. Although these are sep-
arable dimensions, psychometric studies of social judgments
from faces show that these judgments are highly
inter-correlated with each other, with correlations ranging
from 0.60 to 0.80 (Oosterhof and Todorov, 2008; Todorov
et al., 2008a,b). For example, principal components analyses
show that (i) the first component, which indicates general
face valence, accounts for >60% of the variance of judg-
ments; and (ii) trustworthiness and attractiveness judgments
are highly correlated with this valence component. Given
286 SCAN (2013)
these behavioral data, one would expect to observe overlap-
ping regions in neuroimaging studies on attractiveness and
trustworthiness.
For the purposes of this meta-analysis, we focus on studies
on attractiveness and trustworthiness. Typically, such studies
present participants with facial stimuli that vary on the re-
spective dimensioneither systematically manipulated via
computer modeling, or confirmed by independent behavior-
al ratingsand subsequently report brain activity that shows
a linear relationship with changes in facial appearance along
that dimension. For example, some studies have observed
increased responses in the amygdala for untrustworthy
faces (Winston et al., 2002), while other studies have
observed increased responses in the nucleus accumbens
(NAcc) and medial orbitofrontal cortex (mOFC) for attract-
ive faces (Aharon et al., 2001; O’Doherty et al., 2003). More
recent studies have sought to identify regions that show a
quadratic relationship between brain activity and changes in
attractiveness or trustworthiness. Researchers have observed
non-linear responses in the amygdala for both attractive and
unattractive faces (Winston et al., 2007), as well as for both
trustworthy and untrustworthy faces (Todorov et al., 2011).
While there is convergence between the linear and
non-linear approaches, there exists the possibility that
these analyses are tapping distinct processes, wherein areas
that show a linear pattern of activity are coding for face
valence, while areas that show quadratic patterns are
coding something more like face intensity.
The first objective of this article is to systematically ex-
plore the pattern of observed brain activations across pub-
lished neuroimaging studies on face evaluation as a function
of face valence. The second objective is to examine possible
dissociations between linear and non-linear responses. The
third and final objective is to explore potential differences
between trustworthiness and attractiveness studies.
Multilevel kernel density analysis
Meta-analysis is a powerful statistical tool that allows re-
searchers to combine the data sets of a collection of similar
studies to provide a more accurate, robust estimate of the
effect-size of a given phenomena. This approach is wide-
spread within behavioral research, and in recent years,
meta-analyses of neuroimaging studies have become more
common (Fox et al., 1998; Phan et al., 2002; Wager and
Smith, 2003; Wager et al., 2004; Laird et al., 2005; Nielsen
et al., 2005). Meta-analyses of neuroimaging data typically
compute how frequently studies examining a given psycho-
logical phenomenon report activity in a specific brain area
(Kober and Wager, 2010). This approach can be used to
confirm the prevailing thinking regarding what brain areas
are associated with a particular psychological phenomenon
or experience. At the same time, meta-analysis can serve a
more exploratory purposeidentifying regions that are con-
sistently activated across a large number of studies of the
P. Mende-Siedlecki et al.
same psychological phenomenon, but that are not typically
associated with that phenomenon.
Indeed, a meta-analysis of the social evaluation of faces
has been recently published (Bzdok et al., 2011), and in part,
motivated the analyses herein. While we ultimately em-
ployed slightly different selection criteria in choosing studies
to include, we also sought to perform several more targeted
analyses, as noted above. Perhaps more importantly, while
Bzdok and colleagues conducted an activation likelihood es-
timation (ALE) meta-analysis, we use a different statistical
procedure.
Specifically, we use a Multi-level Kernel Density Analysis
(MKDA), which represents an advance in meta-analytic
methods for neuroimaging data, because it accounts for
the fact that individual activation peaks are nested within
contrast maps (maps of particular comparisons within stu-
dies), making these maps the unit of analysis, and not the
peaks (Wager et al., 2008). Further, MKDA models contrast
maps as a random effect, eliminating the possibility of one
contrast dominating the meta-analysis.
We conduct several analyses. First, we analyze activations
across all contrasts showing (i) stronger brain responses to
negativeuntrustworthy and unattractivethan positive
trustworthy and attractivefaces; (ii) stronger responses to
positive than negative faces; and (iii) stronger responses to
positive and negative faces than to neutral faces. Second,
within these contrasts, we also explore potential differences
between trustworthiness and attractiveness studies.
METHODS
Data collection
We searched for neuroimaging studies of the social evaluation
of faces using the online databases PsycINFO and PubMed, as
well as the scholarly article search engine Google Scholar. We
limited our search using combinations of keywords including
‘faces’, ‘social evaluation’, ‘social judgment’, ‘fMRI’, ‘trust-
worthiness’ and ‘attractiveness’. To be included in our
meta-analysis, studies had to involve fMRI or PET investiga-
tions of healthy adults,1 report activations in a standard co-
ordinate systemeither Talairach or Montreal Neurological
Institute (MNI) coordinates, and explicitly state whether
their analyses were performed with fixed or random effects.
With respect to in-scanner tasks, we only included studies in
which subjects either made explicit judgments regarding the
trustworthiness or attractiveness of faces, or were presented
with faces that varied on one of these two dimensions during
an implicit or a passive viewing task, based upon normative
ratings, computer modeling or some other form of categor-
ization. In the case of some studies (Hampshire et al., 2011;
Pochon et al., 2008; Zaki et al., 2011), relevant contrasts were
not originally reported, but were obtained through personal
communication with the respective authors.
1
Pinkham et al. (2008a) and Baas et al. (2008) were neuroimaging studies comparing the social evaluation of
faces in patient populations to healthy controls. When composing our MKDA, we only included the coordinates
yielded from the analysis of the healthy controls’ data.
Social evaluation of faces SCAN (2013) 287

Table 1 Breakdown of Studies Included in the Meta-Analysis

Study Included N Task naturea Study type Valence ROI?c Stimulus category
contrasts
Implicit Explicit Collapsedb Negative Positive Non-Linear

Aharon, et al., 2001 1 6 X attractiveness X extreme

Aron, et al., 2005 1 17 X attractiveness X average
Baas, et al., 2008 4 21 X X X trustworthiness X X average
Blasi, et al., 2009 1 43 X trustworthiness X average
Bray, et al., 2007 2 25 X attractiveness X X extreme
Chatterjee, et al., 2009 4 13 X X attractiveness X X computer generated
Cloutier, et al., 2008 2 48 X attractiveness X X extreme
Engell, et al., 2007 1 15 X trustworthiness X average
Gordon, et al., 2009 2 6 X trustworthiness X X X average
Hampshire, et al., 2011 1 19 X attractiveness X extreme
Iaria, et al., 2008 1 10 X attractiveness X extreme
Kampe, et al., 2001 1 16 X attractiveness X average
Kim, et al., 2007 1 25 X attractiveness X computer generated
Liang, et al., 2010 3 17 X attractiveness X X X X extreme
O’Doherty, et al., 2003 2 25 X attractiveness X X extreme
Pinkham, et al., 2008 1 12 X trustworthiness X X average
Pochon, et al., 2008 1 17 X attractiveness X extreme
Said, et al., 2009 3 31 X trustworthiness X X X average
Said, et al., 2010 1 24 X trustworthiness X computer generated
Smith, et al., 2010 1 26 X attractiveness X average
Todorov, et al., 2008 2 14 X trustworthiness X X X computer generated
Todorov, et al., 2010 2 22 X trustworthiness X X computer generated
Todorov, et al., 2010 2 22 X trustworthiness X X computer generated
Tsukiura, et al., 2010a 3 22 X attractiveness X X X average
Turk, et al., 2004 1 18 X attractiveness X extreme
VanRijn, et al., 2011 3 18 X trustworthiness X X X average
Winston, et al., 2002 2 14 X trustworthiness X X average
Winston, et al., 2007 2 26 X attractiveness X X extreme
Zaki, et al., 2011 1 14 X attractiveness X extreme
a
‘Task nature’ categorizes only the contrasts included in our meta-analysis. In some cases, like Winston et al. (2002), both explicit and implicit paradigms were employed, but only
collapsed analyses were reported.
b
‘Collapsed’ analyses refer to analyses in which neural activity was aggregated across both explicit and implicit tasks.
c
In this column, we note if a given study reported coordinates arising from ROI-based analyses. In some cases, these studies only reported such ROI-based analyses (for instance,
Pinkham et al., 2008). As such, these studies have only been able to impact our supplementary analyses, which incorporate ROI-based analyses in addition to whole-brain
contrasts.

We excluded studies that did not report specific coordin-
ates arising from relevant contrasts, but instead referred to
various ROIs from a functional localizer being more or
less active during specific contrasts (Kranz and Ishai,
2006). In some instances, multiple studies were found
which presented analyses of the same data sets (Todorov
and Engell, 2008; Pinkham et al., 2008b). In these cases,
we only included one study’s reported coordinates, and
this choice was made based upon which version of the
study ultimately presented the more relevant analyses.
Finally, we excluded some studies whose research questions
bordered on ours (for instance, aesthetic judgments of paint-
ings of faces, as in Kawabata and Zeki (2004) or neural re-
sponses to faces similar to the self varying in trustworthiness,
as in Verosky and Todorov (2010) as they ultimately did not
report contrasts that were appropriate for inclusion in our
analyses. These choices are not trivial, as they represent some
of the differences between our meta-analysis and the one
conducted by Bzdok and colleagues (2011), in terms of
study selection.
This search yielded 28 published papers comprising 292
neuroimaging studies on the social evaluation of faces.
Seventeen of these studies were on attractiveness evaluations
and 12 were on trustworthiness or related evaluations (i.e.
‘would you approach or avoid this person’). The latter were
included because such approach/avoidance evaluations
are highly correlated with trustworthiness evaluations
(Todorov, 2008). This set of studies accounted for 52 separ-
ate contrasts (Table 1). For contrasts to be included in our
database, they had to be representative of neural activity that
varied parametrically with either facial attractiveness or
trustworthiness, and furthermore, the direction and linearity
2
We consider a single study to represent an investigation of the neural responses to a given set of stimuli in
the context of one or potentially multiple psychological tasks within the same set of subjects. Thus, Todorov
et al. (2011) represents two separate studies, while Chatterjee et al. (2009)in which the same subjects took
part in explicit and implicit tasks in-scanner on separate scanning daysrepresents one study.
288 SCAN (2013)
of this relationship had to be clearly stated. We excluded
coordinates derived from complex interaction-based ana-
lyses (for instance, stimuli type and gender interactions, as
seen in O’Doherty et al. (2003), as well as coordinates arising
from analyses that were not relevant to our research ques-
tions (e.g. effects of face novelty in Kim et al. (2007).
Further, overlapping contrasts are often reported in the art-
icles surveyed. For instance, Aharon and colleagues (2001)
report separate contrasts detailing neural activity associated
with facial attractiveness for male stimuli, female stimuli,
and collapsed across both kinds of stimuli. In these cases,
we only included the most general reported contrastfor
instance, for Aharon et al. (2001), we used the collapsed
contrast. Studies that report separate results for explicit
and implicit paradigms presented a unique problem (see
Winston et al., 2002; Baas et al., 2008; Chatterjee et al.,
2009). On the one hand, both analyses are relevant to our
main research question, and favoring one paradigm over the
other in these three cases would bias our results in favor of
that task design. On the other hand, the contrasts are un-
deniably non-independent of each other. Ultimately, we
chose to run our analyses using both sets of coordinates
for these three studies, which were entered into our database
as separate contrasts. To confirm that this approach had no
demonstrable impact on our results, we ran complimentary
analyses that only included one contrast per study (i.e. only
the explicit task contrast from the three studies in question).
We observed no practical differences in either the size or
localization of consistent activations.
We tabulated the design particulars and parameters of
each study, as well as the reported activation points for all
relevant contrasts. Specifically, we coded each study in terms
of which coordinate system activations were reported in,
number of participants, whether a fixed or random effects
analysis had been performed, whether activations repre-
sented linear or non-linear effects, whether the task was
explicit or implicit in nature, and whether the reported
activations were the result of a whole-brain or
region-of-interest (ROI) analysis. This coding scheme
served two purposes. Primarily, this information was fed
into the MKDA toolbox and used to determine the proper
weighting scheme for the different studies. Secondarily, it
served as the basis for contrasting studies against each
other on relevant variables. This coding scheme was initially
entered by the first author, with subsequent confirmation
and complete agreement from the second and third authors.
Entered coordinates were checked and re-checked against
their original sources numerous times throughout the
course of setting up our database.
The studies compiled in our database used a variety of face
stimuli. Some studies used computer-generated faces (for
instance, Chatterjee et al., 2009), others used standardized
photograph sets of volunteer subjects (for instance, Engell
et al., 2007), and still others used photographs culled from
magazines and newspapers (for instance, O’Doherty et al.,
P. Mende-Siedlecki et al.
2003). These faces likely differ in terms of their typical-
ityfaces in standardized photographs are more typical
than the extreme faces seen in photographs of models and
actresses. Given recent work suggesting that face typicality
can partially account for the amygdala’s response to the va-
lence of face stimuli (Said et al., 2010), it is possible that
different types of face stimuli (e.g. extremely attractive faces
that are less typical) could lead to different patterns of neural
responses. As such, while we did not exclude studies based
on the sources of the face stimuli, we did keep track of the
source of each study’s stimuli. This allowed for the possibil-
ity of comparing the more typical faces (computer-generated
and standardized sets) against the more atypical faces
(photos of models and actors from print media).
It is important to note that contrasts containing
ROI-based analyses pose a problem for inclusion in
meta-analyses. On the one hand, including coordinates
from ROI-based analyses may bias the results by introducing
researchers’ a priori predictions about which regions are
involved in trustworthiness and attractiveness evaluation.
On the other hand, such analyses represent theoretically
motivated prior research. Further, because some ROIs like
the amygdala and NAcc are relatively small and often diffi-
cult to image, excluding ROI-based analyses may miss im-
portant findings that are consistent across studies. Given
that, we chose to run each of our analyses twiceonce lim-
ited to whole-brain contrasts, and once with ROI-based con-
trasts included. In the interest of space, we chose to report
the whole-brain analyses in the main text, as well as to note
whether or not adding ROI-based contrasts substantially af-
fected the results. (In all cases but one, adding ROI-based
contrasts did not have a substantial effect on analyses. For
the contrast that produced divergent results, we chose to
explicitly note in the text how the two approaches differed.)
The specific results for the ROI-based analyses are reported
in supplemental material. We note that while some studies
reported ROI-based analyses side by side with whole-brain
analyses (for instance, Van Rijn et al., 2011), there are a small
number of studies that reported only ROI-based analyses
(for instance, Pinkham et al., 2008a).
Data analysis
Our MKDA of ‘negative’ contrasts comprised all contrasts in
which brain activity increases as facial stimuli decrease in
either trustworthiness or attractiveness. ‘Positive’ contrasts
comprised all contrasts in which brain activity increases as
facial stimuli increase in either trustworthiness or attractive-
ness. Non-linear, quadratic contrasts comprised all contrasts
in which brain activity increases as facial stimuli increase or
decrease in either trustworthiness or attractiveness relative to
faces at the middle of the continuum. Not all studies
included in our database reported both negative and positive
contrasts. Therefore, neither of our primary MKDAs con-
tains contrasts from every study.
Social evaluation of faces
When performing these analyses, the peak coordinates
from all relevant contrast maps were first separately con-
volved with a 10-mm spherical kernel, yielding comparison
indicator maps (CIMs). Previous meta-analytic work sug-
gests that this is an appropriate default kernel size (Wager
et al. 2003; Salimi-Khorshidi et al., 2009). These CIMs were
subsequently weighted based upon the number of partici-
pants and what type of analysis was performed in each
study, following the same parameters used by Kober and
colleagues (2008). Specifically, each map was first weighted
by the square root of its study’s sample size and subsequently
multiplied by an adjustment factor accounting for the type
of analysis used in the respective study. Random effects stu-
dies were multiplied by an adjustment factor of 1; fixed ef-
fects studies were multiplied by an adjustment factor of .75.
In this fashion, studies received higher weighting if they had
large sample sizes and performed random effects analyses.
Second, the weighted CIMs were averaged together, produ-
cing a density map. Each voxel of this density map attains a
density statistic, P, which is the weighted proportion of con-
trasts included in the MKDA that yield activity within 10 mm
of that voxel. To identify voxels whose P-statistic exceeds the
frequency expected by chance, a Monte Carlo simulation was
conducted. Over 5000 iterations of the Monte Carlo simula-
tion, the observed activation blobs (contiguous regions of ac-
tivation within the CIMs, holding shape constant) from each
CIM were randomly shuffled within a gray-matter mask.
Following each iteration, we recorded both the maximum
whole brain density statistic (P, across all studies) and the
largest cluster of contiguous voxels. These values were used
to create null-hypothesis distributions for the density statistic
and the expected size of clusters, respectively.
Third, the weighted P was subsequently tested against the
resulting null-hypothesis P0 produced by the Monte Carlo
simulation. A similar procedure was used to test for the
significance of the size of the clusters, allowing us to identify
a size threshold at which a certain number of voxels must be
activated contiguously for a given cluster to be deemed sig-
nificant. Hence, we used two types of thresholdsa density
height-based threshold and a cluster size threshold, the latter
derived from a non-parametric cluster-based thresholding
procedure (Nichols and Hayasaka, 2003). For P, the resulting
familywise error rate (FWER)-controlled threshold is the
proportion of studies reporting activity within 10 mm of a
given voxel that exceeds the maximum P-statistic across 95%
of the resulting Monte Carlo maps. These voxels appear on
resulting maps colored in yellow and will be referred to in
our results as exceeding the height-based threshold of the
MKDA. For the cluster size threshold, the resulting
FWER-controlled threshold is the clusters observed at
P < .001 and P < 0.01 whose size exceeds the maximum clus-
ter size across 95% of the Monte Carlo maps. These voxels
appear on resulting maps in orange and pink, respectively,
and will be referred to as exceeding the extent-based thresh-
old. The thresholded maps were overlaid on a canonical MRI
SCAN (2013) 289
image (colin27.img, the single-subject template in SPM5;
http://www.fil.ion.ucl.ac.uk/spm/software/spm5/), which
was co-registered to the MNI brain template.
When reporting areas of consistent activation in our
tables, we provide information on whether each area with-
stood height-based thresholding, extent-based thresholding,
or both. Some areas of activation were sizable enough to pass
extent-based thresholding but not height-based threshold-
ing. Conversely, other areas were highly consistent across
the database and passed height-based thresholding, but
were not sufficiently large to pass extent-based thresholding.
XYZ-coordinates reported in our tables reflect the peak ac-
tivation foci which withstand height-based thresholding, or,
if activations are less consistent, the center of mass of the
cluster at the most stringent level of extent-based threshold-
ing. Further, we report the number of voxels in each cluster
which withstood height-based thresholding, or if activations
are less consistent, the number of voxels at the most strin-
gent level of extent-based thresholding.
We also performed several smaller, more targeted MKDAs
exploring differences between trustworthiness and attractive-
ness studies and performed several additional exploratory
analyses based on stimulus typicality. To perform such ana-
lyses, a simple subtraction yields the relative difference in the
distribution of peaks between the respective contrasts, which
is subsequently thresholded as explained above.
RESULTS
Results across negative contrasts
Eleven studies reported 13 negative contrastswhere brain
activity increased as attractiveness or trustworthiness decrea-
sedacross the whole brain. The MKDA results for these
contrasts are summarized in Table 2 and Figure 1. We
observed highly consistent activation in right amygdala
(withstood height- and extent-thresholding, P < 0.001), as
well as activation in left amygdala that survived extent- but
not height-thresholding (P < 0.001). Four studies reported
ROI-based coordinates for negative contrasts. When we
included these additional coordinates in our analysis as
well, we continued to observe highly consistent activation
in right amygdala (withstood height- and extent-
thresholding, P < 0.001), as well as activations which sur-
vived extent- but not height-thresholding in left amygdala
(P < 0.001), right globus pallidus (P < 0.01) and a large
region of consistent activation encompassing right anterior
insula, right inferior frontal gyrus (IFG) and right ventrolat-
eral prefrontal cortex (vlPFC, P < 0.01, additional results
summarized in Supplementary Table S1).
Results across positive contrasts
Twenty-one studies reported 23 positive contrastswhere
brain activity increased as attractiveness or trustworthiness
increasedacross the whole brain. The MKDA results for
these contrasts are summarized in Table 2 and Figure 2.
We observed highly consistent activation in left caudate
290 SCAN (2013) P. Mende-Siedlecki et al.

Table 2 Areas Consistently Activated During Negative and Positive Face Evaluations
Region Lat x y z Vol %Act

Increased activity for negative evaluations, collapsed across Untrustworthiness and Unattractiveness, whole-brain contrasts only
Basal telencephalon
Amygdala R 20 6 18 1080 0.34 y**
Amygdala L 18 6 18 1328 0.29 **

Increased activity for positive evaluations, collapsed across Trustworthiness and Attractiveness, whole-brain contrasts only
Basal telencephalon
Caudate/nucleus accumbens/medial orbitofrontal cortex L 10 10 4 1888 0.33 y**
Thalamus R 14 16 6 384 0.28 y**
Caudate/right amygdala/anterior insula/inferior frontal gyrus R 4 22 0 76656 0.22 **
Ventral striatum/thalamus/anterior insula/inferior frontal gyrus/ventrolateral prefrontal cortex R 14 2 2 34688 0.20 *

Frontal/insular cortex
Ventromedial prefrontal cortex – 0 42 6 344 0.30 y**
Pregenual cingulate cortex/dorsal anterior cingulate – 2 40 8 96 0.25 y**
Pregenual cingulate cortex – 0 36 2 16 0.25 y**

Note. Stereotactic coordinates representing the areas most consistently activated across full database. We report laterality (Right or Left), XYZ coordinates, number of voxels in
each cluster (Vol), and weighted percentage of CIMs which activated each cluster (%Act).
y, areas withstanding height-based thresholding.
**, areas withstanding extent-based thresholding (p < .001).
*, areas withstanding extent-based thresholding (p < .01).

(unattractive or untrustworthy) and positive-looking (at-

Fig. 1 Consistently activated areas across negative evaluations, showing consistent
right amygdala activation. Yellow voxels withstood height-based thresholding, orange
voxels withstood extent-based thresholding (P<.001).
extending into NAcc and mOFC, right thalamus, vmPFC
and dACC/pgACC (withstood height- and extent-
thresholding, P < 0.001), as well as portions of right amyg-
dala right anterior insula, right IFG (P < 0.001), and bilateral
vlPFC that survived extent- but not height-thresholding (P <
0.01). Four studies reported ROI-based coordinates for
negative contrasts. Including these additional coordinates
in our analysis yielded similar results (summarized in
Supplementary Table S1).
Non-linear responses
Nine studies within our database conducted non-linear
analyses testing for stronger responses to both negative-
tractive or trustworthy) faces than to faces at the middle of
the continuum. Collapsed across both sets of stimuli, we
observed consistent non-linear activation across seven
whole-brain contrasts in the right amygdala extending into
right putamen (withstood height- and extent-thresholding,
P < 0.001, Table 3). Including two additional ROI-based
contrasts in the analysis yielded similar results
(Supplementary Table S2). We note that given the relatively
small number of contrasts documenting non-linear re-
sponses, this analysis is underpowered. Nevertheless, five of
the seven whole-brain contrasts reported activity in right
amygdala.
We also compared non-linear responses against linear re-
sponses, though these comparisons are, by virtue of the
smaller number of non-linear contrasts, unavoidably unba-
lanced. Contrasting negative linear contrasts (13 contrasts)
against non-linear contrasts (seven contrasts), we observed a
ventral portion of the right amygdala that was more consist-
ently active in negative linear contrasts (withstood height-
but not extent-thresholding, Supplementary Table S3), while
a more dorsal portion of the right amygdala was more
consistently active in non-linear contrasts. Including
ROI-based contrasts in the analysis yielded similar results
(Supplementary Table S4). (As this contrast is unbalanced,
we have provided information regarding the frequency of
activation at the peak voxels of those areas that withstood
height-thresholding, Supplementary Table S5A)
Contrasting positive linear contrasts (23 contrasts)
against non-linear contrasts (seven contrasts), we observed
a set of regions that were more consistently active in positive
linear contrasts, including bilateral caudate, vmPFC/OFC,
dACC/pgACC (withstood height- and extent-thresholding,
Social evaluation of faces SCAN (2013) 291

Fig. 2 Consistently activated areas across positive evaluations, including pgACC, vmPFC (A), left caudate/NAcc extending into mOFC (A and B), and right amygdala (C).
Yellow voxels withstood height-based thresholding, orange voxels withstood extent-based thresholding (P < 0.001), and pink voxels withstood extent-based thresholding
(P < 0.01).

Table 3 Consistently Activated Areas Displaying Non-linear Response
Profiles
Region Lat x y z Vol %Act
Non-Linear responses, collapsed across Attractiveness and Trustworthiness studies
Basal telencephalon
Amygdala R 20 2 10 1040 0.63 y
Note. Stereotactic coordinates representing the areas most consistently activated
across full database. We report laterality (Right or Left), XYZ coordinates, number
of voxels in each cluster (Vol), and weighted percentage of CIMs which activated each
cluster (%Act).
y, areas withstanding height-based thresholding.
P < 0.001), right thalamus (withstood height-thresholding,
P < 0.001), sgACC and bilateral vlPFC (P < 0.01,
Supplementary Table S3). Conversely, we observed a
region of right dorsal amygdala extending into right puta-
men (withstood height- but not extent-thresholding, results
summarized in Supplementary Table S3) that was more con-
sistently active in non-linear contrasts. Including ROI coord-
inates in the analysis yielded similar results (noted in
Supplementary Table S4). (Information regarding the fre-
quency of activation at the peak voxels of those areas that
withstood height-thresholding is provided in Supplementary
Table S5B.)
Negative linear responses in attractiveness and
trustworthiness studies
We contrasted negative linear responses in attractiveness (six
contrasts) and trustworthiness studies (seven contrasts),
observing one activation in the right amygdala that was
more consistently active for negative linear responses to
trustworthiness than attractiveness (withstood height- and
extent-thresholding, P < 0.001). We observed no regions
that were consistently more active for negative linear re-
sponses to attractiveness than trustworthiness. (Results are
summarized in Supplementary Table S5.) Including ROI co-
ordinates (from one unattractiveness study and three un-
trustworthiness studies) in the analysis yielded similar
results (noted in Supplementary Table S6).
Positive linear responses in attractiveness and
trustworthiness studies
We contrasted positive linear responses in attractiveness (18
contrasts) and trustworthiness (5 contrasts). We observed
activations in left caudate extending into NAcc, vmPFC/
OFC and pgACC extending dorsally into dACC (withstood
height- and extent-thresholding, P < 0.01) that were more
consistent for positive linear responses to attractiveness
than trustworthiness. We observed no regions that were
292 SCAN (2013) P. Mende-Siedlecki et al.

Fig. 3 Consistently activated areas in the contrast ‘Extreme’ Attractiveness > Trustworthiness, (A) NAcc extending into mOFC, (B) pgACC and vmPFC. (The Average
Attractiveness > Trustworthiness contrast produces no regions of consistent activation.) Yellow voxels withstood height-based thresholding and pink voxels withstood
extent-based thresholding (P < 0.01).

consistently more active for positive linear responses to
trustworthiness than attractiveness. (Results are summarized
in Supplementary Table S5.) Including ROI coordinates
(from one attractiveness study and three trustworthiness
studies) in the analysis yielded similar results (noted in
Supplementary Table S6). (Information regarding the fre-
quency of activation at the peak voxels of those areas that
withstood height-thresholding is provided in Supplementary
Table S5C.)
Separating attractiveness studies by stimulus type
The differences between trustworthiness and attractiveness
studies are interesting but also puzzling given that evalu-
ations on these two dimensions are highly correlated.
There were no obvious differences between these two sets
of studies (for instance, they were well-balanced be-
tween implicit and explicit tasks) except for the nature of
the face stimuli used in the studies. Whereas eleven of the
attractiveness studies used atypical, extremely attractive
faces (culled from magazines and print media, often of
models), none of the trustworthiness studies used such
faces (typically, these were standardized sets of faces or
computer-generated faces).
If the differences between attractiveness and trustworthi-
ness studies are partly due to differences in stimuli, then the
regions that differentiate these studies should also appear in
contrasts involving the extremeness of faces. We can test this
proposition by splitting attractiveness studies into two
groupsthose that used extremely attractive stimuli and
those that used average or computer-generated stimuli.
Comparing extreme attractiveness studies to the set of trust-
worthiness studies should yield areas of consistent activation
in NAcc/caudate and mOFC, for example, while there should
be fewer differences between average attractiveness and trust-
worthiness studies.
Indeed, when contrasting extreme attractiveness (11 con-
trasts) against trustworthiness (8 contrasts), we observed
consistent activation in left caudate and NAcc, extending
into mOFC, pgACC, and vmPFC (withstood height- and
extent-thresholding, p < .01; see Figure 3, results summarized
in Supplemental Table 7). Further, we observed a consistent
pattern of activation centered in pgACC and extending
broadly into both vmPFC and vlPFC that withstood
extent-thresholding (p < .01) but not height-thresholding.
Contrasting average attractiveness against trustworthiness
produced no areas of consistent activation.
Similarly within the set of attractiveness studies, when
contrasting studies that used extremely attractive faces (11
contrasts) against studies that used more typical faces (7
contrasts), we observed consistent activation in left caudate,
vmPFC/mOFC, and pgACC/dACC (withstood height- and
extent-thesholding, p < .01), while a larger activation
extending broadly through mOFC, vmPFC and vlPFC with-
stood extent-thresholding (p < .01), but not height-
thresholding. The reverse contrast produced no areas of con-
sistent activation.
Further, differences due to face stimuli should be appar-
ent in studies that used implicit evaluation paradigms.
Because no evaluative dimension is specified in such para-
digms, stimulus properties should drive the neural responses.
Contrasting implicit paradigm studies that used extremely
attractive faces with implicit paradigm studies that used
more typical faces produced consistent activation in right
amygdala, left caudate extending into NAcc and right inferior
frontal gyrus (withstood height- but not extent thresholding;
results are summarized in Supplemental Table S7).
DISCUSSION
Using multi-level kernel density analysis, a statistically rigor-
ous method of meta-analysis that treats contrasts as the unit
Social evaluation of faces
of analysis instead of individual activation peaks, we per-
formed a meta-analysis on 29 neuroimaging studies of the
social evaluation of faces. We split these studies by valence
into two MKDAs, one focusing on brain responses to nega-
tive evaluations like unattractiveness and untrustworthiness,
and the other focusing on brain responses to positive evalu-
ations like attractiveness and trustworthiness.
Our negative MKDA revealed the most consistent activa-
tion in right amygdala. Less consistent areas of activation
were observed in left amygdala, right anterior insula, right
IFG, right vlPFC and right globus pallidus. These results are
remarkably consistent with previous findings regarding the
neural responses to angry faces (Morris et al., 1998; Whalen
et al., 2001; Monk et al., 2006; Dannlowski et al., 2007).
Amygdala responses to angry faces have been widely
observed and characterized (Morris et al., 1998; Whalen
et al., 2001; Nomura et al., 2004; Taylor et al., 2006;
Dannlowski et al., 2007; Monk et al., 2008; Vrticka et al.,
2008). Furthermore, a functional connectivity between the
amygdala and vlPFC has been proposed and demonstrated
(Nomura et al., 2004; Taylor et al., 2006; Monk et al., 2006,
2008), suggesting that in response to angry faces, the vlPFC
may serve to modulate amygdala reactivity, effectively reg-
ulating emotional responses. Right IFG (Dannlowski et al.,
2007), right insula (Dannlowski et al., 2007; Vrticka et al.,
2008) and right globus pallidus (Jackson et al., 2008) have
also all been implicated in the neural response to angry faces.
Our positive MKDA revealed highly consistent activations
in left caudate extending into NAcc/mOFC, vmPFC, dACC/
pgACC, right thalamus as well as less consistent activations
in right amygdala, insula, IFG and vlPFC. Once again, this
pattern of activations bears a strong resemblance to that
associated with a different emotional expressionhappiness.
Happy faces have been observed to elicit responses in parts
of the striatum (Morris et al., 1996; Phillips et al., 1998;
Fu et al. 2007; Vrticka et al., 2008), mPFC (Phillips et al.,
1998; Kesler-West et al., 2001), ACC (Dolan et al., 1996;
Phillips et al., 1998; Kesler-West et al., 2001; Vrticka et al.,
2008), thalamus (Dolan et al., 1996), IFG (Dolan et al., 1996)
and insula (Lee et al., 2002). Furthermore, a number of
studies have observed amygdala responses to happy faces
(Breiter et al., 1996; Canli et al., 2002; Pessoa et al., 2002;
Yang et al., 2002; Somerville et al., 2004).
The consistent activation in the left caudate nucleus, ex-
tending broadly into the nucleus accumbens, suggests that
positive evaluation of faces may depend, in part, on the re-
cruitment of structures implicated in reward-processing
(Knutson et al., 2001a,b; Haruno et al., 2004). However,
we note that consistent activation in this area was almost
entirely driven by attractiveness contrasts, and, therefore,
may not be part of a general network for face evaluation.
Nonetheless, the highly consistent presence of these areas in
our meta-analysis suggests that under certain task and
stimulus conditions, attractive faces modulate activity in
reward-related regions of the brain.
SCAN (2013) 293
The similarities between the neural correlates of negatively
and positively evaluated faces and angry and happy faces,
respectively, parallels perceptual similarities between these
types of faces. In computer models of facial trustworthiness,
extreme untrustworthiness resembles anger and extreme
trustworthiness resembles happiness (Oosterhof and
Todorov, 2008, 2009; Todorov et al., 2008a,b). Further, be-
havioral adaptation studies suggest common neural under-
pinnings for evaluations of trustworthiness and anger/
happiness (Engell et al., 2010). These observations are con-
sistent with the emotion overgeneralization hypothesis
(Montepare and Dobish, 2003; Todorov et al., 2008a,b;
Zebrowitz and Montepare, 2008; Said et al., 2009a,b), ac-
cording to which evaluative judgments of faces are based
on configurations of facial features resembling emotional
expressions. In the context of positive and negative evalu-
ation, these configurations signal approach and avoidance
behaviors, respectively (Todorov, 2008). Our meta-analysis
findings are also consistent with the hypothesis that novel
faces are automatically evaluated with respect to their ap-
proach/avoidance value.
The role of the amygdala in face evaluation
The amygdala is critical for adaptive social behavior
(Adolphs, 2010; Sander et al., 2003) and, possibly, for
normal face perception and evaluation (Todorov, 2011).
Large meta-analyses of PET and fMRI studies on emotional
processing show that faces are one of the most potent stimuli
for eliciting responses in the amygdala (Costafreda et al.,
2008; Sergerie et al., 2008). The role of the amygdala in
face evaluation is also consistent with neurophysiology find-
ings of face selective responses in the amygdala (Nakamura
et al., 1992; Rolls, 2000a,b; Gothard et al., 2007). The
amygdala receives input from the inferior temporal (IT)
cortex and projects back not only to IT cortex but also to
extrastriate and striate visual areas (Amaral et al., 1992). The
amygdala also has strong interconnections with rACC, OFC,
mPFC, basal ganglia and anterior insula. This anatomical
position of the amygdala allows for it to serve as an affective
hub of information. The current findings, together with the
findings of a recent ALE-based meta-analysis of a smaller
and only partially overlapping set of 16 studies on face evalu-
ation (Bzdok et al., 2011), further buttress the importance of
the amygdala in face perception and evaluation.
Importantly, the amygdala responded not only to nega-
tively evaluated faces but also to positively evaluated faces,
consistent with meta-analyses of its responses to the valence
of emotional expressions (Sergerie et al., 2008). Interestingly,
we observed different loci of activation within the amygdala
for linear and non-linear responses (Figure 4). Whereas a
ventral portion responded more consistently to negative
faces only, a dorsal portion of the amygdala responded
more consistently to both negative and positive faces than
to neutral faces. This dissociation of linear and non-linear
responses in the human amygdala parallels the findings of a
294 SCAN (2013)
Fig. 4 Linear and non-linear response patterns in right amygdala. Blue indicates voxels more consistently active across non-linear contrasts, red indicates voxels more
consistently active across negative evaluations, and green indicates voxels consistently active across positive evaluations. Blue and red clusters withstood height-based
thresholding, while the green cluster withstood extent-based thresholding (P < 0.001).
high-resolution fMRI study on non-human primates
(Hoffman et al., 2007). Hoffman and colleagues observed a
linear response in ventral portions of the amygdala (com-
prising the basolateral amygdala)specifically, stronger
responses to threatening faces and progressively weaker re-
sponses to neutral and appeasing faces. However, in a more
dorsal portion (comprising the central nucleus and the bed nu-
cleus of the stria terminalis), they observed a non-linear re-
sponsestronger responses to both threatening and appeasing
faces than to neutral faces. This ventral/dorsal distinction also
parallels a distinction made by Whalen and his colleagues
(Whalen et al., 2001; Kim et al., 2003, Somerville, et al.,
2006; Davis, et al., 2010). They have argued that while the
ventral portion of the amygdala is involved in processing va-
lence, the dorsal portion of the amygdala is recruited in deter-
mining the value of ambiguous information (e.g. expressions
of surprise) in a given context. These authors suggest further
that given the dorsal amygdala’s response to surprised (Kim
et al., 2003), fearful (Whalen et al., 1998, 2001) and happy faces
(Breiter et al., 1996; Whalen et al., 1998), it may be tracking the
salience of these faces, more generally. This hypothesis is con-
sistent with the current findings.
These findings open the door to future work along those
lines. One possibility is that there exist separate populations
of neurons within the amygdala that code for stimulus va-
lence and stimulus salience, respectively. Ultimately, the
findings are in line with previous work proposing a shift
away from conceptualizing the amygdala as simply a fear
or threat module and instead toward an account of the
amygdala as also tracking stimulus intensity (Anderson
et al., 2003; Small et al., 2003) or motivational salience
(Sander et al., 2003; Cunningham et al., 2008; Adolphs,
2010; Todorov, 2011). These findings also serve as an excel-
lent reminder that one of the additional benefits of the
meta-analytic method is the possibility of generating new,
testable hypotheses for future research.
Faces that are tagged as affectively significant in the amyg-
dala can be further processed in prefrontal regions, which, in
P. Mende-Siedlecki et al.
turn, can serve to modulate amygdala activity. Prefrontal-
amygdala connections have been explored in the vmPFC
(Quirk et al., 2003, Heinz et al., 2004), as well as
the pgACC (Pezawas et al., 2005; Stein et al., 2007;
Zink et al., 2010), both of which were observed as consist-
ently activated across our set of positive contrasts.
Stimulus effects on neuroimaging findings
We also performed several smaller MKDAs to compare be-
tween study type, within negative and positive linear re-
sponses. These more targeted MKDAs offered evidence
that our negative and positive analyses were driven by un-
trustworthiness and attractiveness, respectively. These two
sets of studies were associated with different loci of activa-
tions: the right amygdala was more consistently active as
facial trustworthiness decreased, while the NAcc/caudate
and vmPFC/pgACC were all more consistently active as
facial attractiveness increased. This distinction mirrors the
results we observed in our primary analyses. In contrast, no
brain regions were consistently activated across contrasts
where facial attractiveness decreased or facial trustworthiness
increased, respectively, even when including ROI coordin-
ates in the analyses. We should note that these contrastses-
pecially the comparison between positive linear responses in
attractiveness and trustworthiness studiesare certainly
unbalanced, rendering the results more suggestive than
confirmative.
As noted in the introduction, given that attractiveness and
trustworthiness judgments from faces are highly correlated,
this pattern is puzzling. These differences between trust-
worthiness and attractiveness studies cannot be explained
by researchers’ a priori focus on different regions because
our results hold even for whole brain analyses that did not
include ROI coordinates from individual studies.
The apparent differences in the neural bases of attractive-
ness and trustworthiness are also puzzling in the context of
studies that used the same set of faces to examine responses
to facial attractiveness and trustworthiness (Todorov and
Social evaluation of faces
Fig. 5 The relationship between perceptions of face typicality and face attractiveness for (A) a standardized set of faces (the Karolinska set) and (B) faces sampled to be
extremely attractive from websites of models. The judgments are shown in standardized units. Each point of the plots represents a face.
Engell, 2008). Specifically, Todorov and Engell re-analyzed
the data from Engell et al. (2007), using 14 different social
judgments of the same set of faces. Most of the brain re-
sponses were accounted by a general valence dimension
rather than by specific dimensions such as attractiveness
and trustworthiness (both of these were highly correlated
with this dimension).
What could be driving the differences in neuroimaging
studies on attractiveness and trustworthiness? One possibil-
ity is that the type of faces used in these studies may lead to
different responses. Specifically, a third variable that is cor-
related with both trustworthiness and attractiveness but
could vary across sets of faces may account for such differ-
ences. One candidate is face typicality. Recently, Said and
colleagues (2010) showed that coding face typicality is a
more parsimonious explanation of prior findings of the in-
volvement of the amygdala in face evaluation than coding
face valence.
Face typicality could vary across data sets and lead to
different results. For example, in many standardized data
sets of natural faces, typicality is positively correlated with
both attractiveness and trustworthiness judgments
(Figure 5A). In studies using these stimuli, the amygdala
shows stronger responses to more atypical faces that hap-
pened also to be more negative (Todorov and Engell, 2008).
In studies using artificial stimuli created by a statistical
model, the most atypical faces are faces at the extremes of
the dimension. In such studies, the amygdala responds to
more atypical faces that happened to be more positive or
more negative (Said et al., 2010). The important distinction
here is not between real and artificial faces. Judgments of
artificial faces that have not been manipulated to exaggerate
differences along social dimensions are linearly correlated
with their perceived typicality. Finally, in attractiveness stu-
dies that use extremely attractive faces (e.g. Aharon et al.,
2001), the most attractive faces may be the least typical
SCAN (2013) 295
(Figure 5B). In such studies, the amygdala may respond to
both extremely attractive and extremely unattractive faces as
observed in Winston et al. (2007).
The typicality hypothesis predicts that faces that system-
atically differ in their perceived typicality may lead to
different neural responses. In fact, in the contrast of attract-
iveness studies using extreme faces and studies using more
typical faces, focusing specifically on studies employing im-
plicit paradigms, we observed consistent patterns of activa-
tion in right amygdala and NAcc/caudate. This suggests that
when task demands are controlled for, the driving force
behind NAcc and caudate activations observed in these
studies was the usage of extremely attractive, atypical faces.
This result also lends additional support to the suggestion
that extreme, atypical faces will drive amygdala activity,
regardless of their trustworthiness. Further, the consistent
activations in vmPFC and pgACC that were observed
across attractiveness contrasts but not trustworthiness con-
trasts can also be accounted for by face typicality.
Contrasting extreme attractiveness and trustworthiness con-
tinued to produce consistent activation in these regions,
while contrasting more typical attractiveness against trust-
worthiness did not.
It may be the case that in the context of face evaluation,
some of the regions implicated in reward processing are only
activated upon the presence of real and extremely attractive
faces, or the goal to evaluate face attractiveness, or some
combination of stimulus features and task demands.
Unfortunately, we do not have a sufficient number of studies
to test for more specific effects.
Recommendations for studies using face stimuli
Our findings suggest that the type of face stimuli selected for
a particular study matters a great deal. For example, using
more ‘extreme’ faces resulted in more consistently observed
activation in the NAcc. Given that stimuli are often selected
296 SCAN (2013)
in an ad hoc fashion and rarely shared among research
groups, this complicates comparisons across studies.
Moreover, it undermines the generalizability of results.
To overcome these problems, researchers need to use a
shared set of stimuli, not necessarily the same stimuli but
stimuli sampled from a common pool. One approach is to
use parametrically manipulated faces generated by an expli-
citly specified statistical model (e.g. Oosterhof and Todorov,
2008; Todorov and Oosterhof, 2011). This approach has the
benefit of providing researchers with a full spectrum of
facesone that is not biased towards one portion of a
given dimension. Our laboratory has made a number of
such databases available for academic research (http://web
script.princeton.edu/tlab/databases/). However, artificial
faces may not be the best stimuli for many investigators.
In this case, it would be best to create a common bank of
stimuli that are shared with other research groups. These
stimuli could be validated on a number of important vari-
ables such as typicality, and these variables could be further
used to facilitate comparisons across studies.
SUPPLEMENTARY DATA
Supplementary data are available at SCAN online.
Conflict of Interest
None declared.
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